Pteranodon was not a dinosaur. By definition, all dinosaurs belong to the groups Saurischia and Ornithischia, which exclude pterosaurs. Nonetheless, Pteranodon is frequently featured in dinosaur books and is strongly associated with dinosaurs by the general public.[2]

Pteranodon pieces are extremely well represented in the fossil record, allowing for detailed descriptions of their anatomy and analysis of their life history. Over 1,000 specimens have been identified, though less than half are complete enough to give researchers good information on the anatomy of the animal. Still, this is more fossil material than is known for any other pterosaur, and it includes both male and female specimens of various age groups and, possibly, species.[2]

Size of P. longiceps male (green) and female (orange) compared with a human

Adult Pteranodon specimens from the two major species can be divided into two distinct size classes. The smaller class of specimens have small, rounded head crests and very wide pelvic canals, even wider than those of the much larger size class. The size of the pelvic canal probably allowed the laying of eggs, indicating that these smaller adults are females. The larger size class, representing male individuals, have narrow hips and very large crests, which were probably for display.

Adult male Pteranodon were among the largest pterosaurs, and were the largest flying animals known until the late 20th century, when the giant azhdarchid pterosaurs were discovered.[2] The wingspan of an average adult male Pteranodon was 5.6 metres (18 ft). Adult females were much smaller, averaging 3.8 metres (12 ft) in wingspan. The largest ' from the Niobrara Formation measured 6.25 metres (20.5 ft) from wingtip to wingtip. An even larger specimen is known from the Pierre Shale Formation, with a wingspan of 7.25 metres (23.8 ft), though this specimen may belong to the distinct genus and species Geosternbergia maysei.[2] While most specimens are found crushed, enough fossils exist to put together a detailed description of the animal.

Methods used to estimate the weight of large male Pteranodon specimens (those with wingspans of about 7 meters) have been notoriously unreliable, producing a wide range of estimates from as low as 20 kilograms (44 lb) and as high as 93 kilograms (205 lb). In a review of pterosaur size estimates published in 2010, researchers Mark Witton and Mike Habib demonstrated that the latter, largest estimates are almost certainly incorrect given the total volume of a Pteranodon body, and could only be correct if the animal "was principally comprised of aluminium."[3] Witton and Habib considered the methods used by researchers who obtained smaller weight estimates equally flawed. Most have been produced by scaling modern animals such as bats and birds up to Pteranodon size, despite the fact that pterosaurs have vastly different body proportions and soft tissue anatomy from any living animal.[3]

Unlike earlier pterosaurs such as Rhamphorhynchus and Pterodactylus, Pteranodon had toothless beaks, similar to those of birds. Pteranodon beaks were made of solid, bony margins that projected from the base of the jaws. The beaks were long, slender, and ended in thin, sharp points. The upper jaw was longer than the lower jaw. The upper jaw was curved upward; while this normally has been attributed only to the upward-curving beak, one specimen (UALVP 24238) has a curvature corresponding with the beak widening towards the tip. While the tip of the beak is not known in this specimen, the level of curvature suggests it would have been extremely long. The unique form of the beak in this specimen led Alexander Kellner to assign it to a distinct genus, Dawndraco, in 2010.[4]

The most distinctive characteristic of Pteranodon is its cranial crest. These crests consisted of skull bones (frontals) projecting upward and backward from the skull. The size and shape of these crests varied due to a number of factors, including age, sex, and species. Male Pteranodon sternbergi, the older species of the two described to date (and sometimes placed in the distinct genus Geosternbergia), had a more vertical crest with a broad forward projection, while their descendants, Pteranodon longiceps, evolved a narrower, more backward-projecting crest.[1] Females of both species were smaller and bore small, rounded crests.[5] The crests were probably mainly display structures, though they may have had other functions as well.[6]

Other distinguishing characteristics that set Pteranodon apart from other pterosaurs include narrow neural spines on the vertebrae, plate-like bony ligaments strengthening the vertebrae above the hip, and a relatively short tail in which the last few vertebrae are fused into a long rod.[7] The entire length of the tail was about 3.5% as long as the wingspan, or up to 25 centimetres (9.8 in) in the largest males.[7]

Pteranodon fossils are known primarily from the Niobrara Formation of the central United States. Broadly defined, Pteranodon existed for more than four million years, during the late Coniacian to early Campanian stages of the Cretaceous period.[5] The genus is present in most layers of the Niobrarra Formation except for the upper two; in 2003, Kenneth Carpenter surveyed the distribution and dating of fossils in this formation, demonstrating that Pteranodon sternbergi existed there from 88 to 85 million years ago, while P. longiceps existed between 86 and 84.5 million years ago. A possible third species, which Kellner named Geosternbergia maysei in 2010, is known from the Sharon Springs member of the Pierre Shale Formation in Kansas, Wyoming, and South Dakota, dating to between 81.5 and 80.5 million years ago.[8]

In the early 1990s, Bennett noted that the two major morphs of pteranodont present in the Niobrara Formation were precisely separated in time with little, if any, overlap. Due to this, and to their gross overall similarity, he suggested that they probably represent chronospecies within a single evolutionary lineage lasting about 4 million years. In other words, only one species of Pteranodon would have been present at any one time, and P. sternbergi (or Geosternbergia) in all likelihood was the direct ancestor species of P. longiceps.[2]

Map of North America during the mid-Cretaceous Period, illustrating the Western Interior Seaway

Specimens assigned to Pteranodon have been found in both the Smoky Hill Chalk deposits of the Niobrara Formation, and the slightly younger Sharon Springs deposits of the Pierre Shale Formation. When Pteranodon was alive, this area was covered by a large inland sea, known as the Western Interior Seaway. Famous for fossils collected since 1870, these formations extend from as far south as Kansas in the United States to Manitoba in Canada. However, Pteranodon specimens (or any pterosaur specimens) have only been found in the southern half of the formation, in Kansas, Wyoming, and South Dakota. Despite the fact that numerous fossils have been found in the contemporary parts of the formation in Canada, no pterosaur specimens have ever been found there. This strongly suggests that the natural geographic range of Pteranodon covered only the southern part of the Niobrara, and that its habitat did not extend farther north than South Dakota.[5]

Some very fragmentary fossils belonging to pteranodontian pterosaurs, and possibly Pteranodon itself, have also been found on the Gulf Coast and East Coast of the United States. For example, some bone fragments from the Mooreville Formation of Alabama and the Merchantville Formation of Delaware may have come from Pteranodon, though they are too incomplete to make a definite identification.[5] Some remains from Japan have also been tentatively attributed to Pteranodon, but their distance from its known Western Interior Seaway habitat makes this identification unlikely.[5]

Life restoration of an adult male P. longiceps in flight

Pteranodon longiceps would have shared the sky with the giant-crested pterosaur Nyctosaurus. Compared to P. longiceps, which was a very common species, Nyctosaurus was rare, making up only 3% of pterosaur fossils from the formation. Also less common was the early toothed bird, Ichthyornis.[8]

It is likely that, as in other polygynous animals (in which males compete for association with harems of females), Pteranodon lived primarily on offshore rookeries, where they could nest away from land-based predators and feed far from shore; most Pteranodon fossils are found in locations which at the time, were hundreds of kilometres from the coastline.[6]

Fossils from terrestrial dinosaurs also have been found in the Niobrara Chalk, suggesting that animals who died on shore must have been washed out to sea (one specimen of a hadrosaur appears to have been scavenged by a shark).[10]

The wing shape of Pteranodon suggests that it would have flown rather like a modern-day albatross. This is based on the fact that Pteranodon had a high aspect ratio (wingspan to chord length) similar to that of the albatross — 9:1 for Pteranodon, compared to 8:1 for an albatross. Albatrosses spend long stretches of time at sea fishing, and use a flight pattern called "dynamic soaring" which exploits the vertical gradient of wind speed near the ocean surface to travel long distances without flapping, and without the aid of thermals (which do not occur over the open ocean the same way they do over land).[11] While most of a Pteranodon flight would have depended on soaring, like long-winged seabirds, it probably required an occasional active, rapid burst of flapping, and studies of Pteranodon wing loading (the strength of the wings vs. the weight of the body) indicate that they were capable of substantial flapping flight, contrary to some earlier suggestions that they were so big they could only glide.[3]

Like other pterosaurs, Pteranodon probably took off from a standing, quadrupedal position. Using their long forelimbs for leverage, they would have vaulted themselves into the air in a rapid leap. Almost all of the energy would have been generated by the forelimbs. The upstroke of the wings would have occurred when the animal cleared the ground followed by a rapid down-stroke to generate additional lift and complete the launch into the air.[3]

The diet of Pteranodon is known to have included fish; fossilized fish bones have been found in the stomach area of one Pteranodon, and a fossilized fish bolus has been found between the jaws of another Pteranodon, specimen AMNH 5098. Numerous other specimens also preserve fragments of fish scales and vertebrae near the torso, indicating that fish made up a majority of the diet of Pteranodon (though they may also have taken invertebrates).[2]

Traditionally, most researchers have suggested that Pteranodon would have taken fish by dipping their beaks into the water while in low, soaring flight. However, this was probably based on the assumption that the animals could not take off from the water surface.[2] It is more likely that Pteranodon could take off from the water, and would have dipped for fish while swimming rather than while flying. Even a small, female Pteranodon could have reached a depth of at least 80 centimetres (31 in) with its long bill and neck while floating on the surface, and they may have reached even greater depths by plunge-diving into the water from the air like some modern long-winged seabirds.[2] In 1994, Bennett noted that the head, neck, and shoulders of Pteranodon were as heavily built as diving birds, and suggested that they could dive by folding back their wings like the modern Gannet.[2]

Pteranodon was notable for its skull crest, though the function of this crest has been a subject of debate. Most explanations have focused on the blade-like, backward pointed crest of male P. longiceps, however, and ignored the wide range of variation across age and sex. The fact that the crests vary so much rules out most practical functions other than for use in mating displays.[12] Therefore, display was probably the main function of the crest, and any other functions were secondary.[6]

Scientific interpretations of the crest's function began in 1910, when George Francis Eaton proposed two possibilities: an aerodynamic counterbalance and a muscle attachment point. He suggested that the crest might have anchored large, long jaw muscles, but admitted that this function alone could not explain the large size of some crests.[13] Bennett (1992) agreed with Eaton's own assessment that the crest was too large and variable to have been a muscle attachment site.[6] Eaton had suggested that a secondary function of the crest might have been as a counterbalance against the long beak, reducing the need for heavy neck muscles to control the orientation of the head.[13] Wind tunnel tests showed that the crest did function as an effective counterbalance to a degree, but Bennett noted that again, the hypothesis focuses only on the long crests of male P. longiceps, not on the larger crests of P. sternbergi and very small crests that existed among the females. Bennett found that the crests of females had no counterbalancing effect, and that the crests of male P. sternbergi would, by themselves, have a negative effect on the balance of the head. In fact, side to side movement of the crests would have required more, not less, neck musculature to control balance.[6]

Short crested P. longiceps holotype specimen YPM1177, now interpreted as a female individual

In 1943, Dominik von Kripp suggested that the crest may have served as a rudder, an idea embraced by several later researchers.[6][14] One researcher, Ross S. Stein, even suggested that the crest may have supported a membrane of skin connecting the backward-pointing crest to the neck and back, increasing its surface area and effectiveness as a rudder.[15] The rudder hypothesis again, does not take into account females nor P. sternbergi, which had an upward-pointing, not backward-pointing crest. Bennett also found that even in its capacity as a rudder, the crest would not provide nearly so much directional force as simply maneuvering the wings. The suggestion that the crest was an air brake, and that the animals would turn their heads to the side in order to slow down, suffers from a similar problem.[16] Additionally, the rudder and air brake hypotheses do not explain why such large variation exists in crest size even among adults.[6]

Alexander Kellner suggested that the large crests of the pterosaur Tapejara, as well as other species, might be used for heat exchange, allowing these pterosaurs to absorb or shed heat and regulate body temperature, which also would account for the correlation between crest size and body size. There is no evidence of extra blood vessels in the crest for this purpose, however, and the large, membranous wings filled with blood vessels would have served that purpose much more effectively.[6]

With these hypotheses ruled out, the best-supported hypothesis for crest function seems to be as a sexual display. This is consistent with the size variation seen in fossil specimens, where females and juveniles have small crests and males large, elaborate, variable crests.[6]

Adult Pteranodon specimens may be divided into two distinct size classes, small and large, with the large size class being about one and a half times larger than the small class, and the small class being twice as common as the large class. Both size classes lived along side each other, and while researchers had previously suggested that they represent different species, Christopher Bennett showed that the differences between them are consistent with the concept that they represent females and males, and that Pteranodon species were sexually dimorphic. Skulls from the larger size class preserve large, upward and backward pointing crests, while the crests of the smaller size class are small and triangular. Some larger skulls also show evidence of a second crest that extended long and low, toward the tip of the beak, which is not seen in smaller specimens.[6]

The sex of the different size classes was determined, not from the skulls, but from the pelvic bones. Contrary to what may be expected, the smaller size class had disproportionately large and wide-set pelvic bones. Bennett interpreted this as indicating a more spacious birth canal, through which eggs would pass. He concluded that the small size class with small, triangular crests represent females, and the larger, large-crested specimens represent males.[6]

Note that the overall size and crest size also corresponds to age. Immature specimens are known from both females and males, and immature males often have small crests similar to adult females. Therefore, it seems that the large crests only developed in males when they reached their large, adult size, making the sex of immature specimens difficult to establish from partial remains.[17]

The fact that females appear to have outnumbered males two to one suggests that, as with modern animals with size-related sexual dimorphism, such as sea lions and other pinnipeds, Pteranodon might have been polygynous, with a few males competing for association with groups consisting of large numbers of females. Similar to modern pinnipeds, Pteranodon may have competed to establish territory on rocky, offshore rookeries, with the largest, and largest-crested, males gaining the most territory and having more success mating with females. The crests of male Pteranodon would not have been used in competition, but rather as "visual dominance-rank symbols", with display rituals taking the place of physical competition with other males. If this hypothesis is correct, it also is likely that male Pteranodon played little to no part in rearing the young; such a behavior is not found in the males of modern polygynous animals who father many offspring at the same time.[6]

Historically, the terrestrial locomotion of Pteranodon, especially whether it was bipedal or quadrupedal, has been the subject of debate. Today, most pterosaur researchers agree that pterosaurs were quadrupedal, thanks largely to the discovery of pterosaur trackways.[18]

The possibility of aquatic locomotion via swimming has been discussed briefly in several papers (Bennett 2001, 1994, and Bramwell & Whitfield 1974).

Pteranodon was the first pterosaur found outside of Europe. Its fossils first were found by Othniel Charles Marsh in 1870, in the Late Cretaceous Smoky Hill Chalk deposits of western Kansas. These chalk beds were deposited at the bottom of what was once the Western Interior Seaway, a large shallow sea over what now is the midsection of the North American continent. These first specimens, YPM 1160 and YPM 1161, consisted of partial wing bones, as well as a tooth from the prehistoric fish Xiphactinus, which Marsh mistakenly believed to belong to this new pterosaur (all known pterosaurs up to that point had teeth). In 1871, Marsh named the find Pterodactylus Oweni, assigning it to the well-known (but much smaller) European genusPterodactylus. Marsh also collected more wing bones of the large pterosaur in 1871. Realizing that the name Pterodactylus oweni already had been used in 1864 for a specimen of the European Pterodactylus, Marsh re-named his North American pterosaur Pterodactylus occidentalis, meaning "Western wing finger," in his 1872 description of the new specimen. He also named two additional species, based on size differences: Pterodactylus ingens (the largest specimen so far), and Pterodactylus velox (the smallest).[5]

Early illustration of Ornithochirus umbrosus (now Pteranodon), when teeth erroneously were attributed to the species and the crest was unknown, 1872

Meanwhile, Marsh's rival Edward Drinker Cope also had unearthed several specimens of the large North American pterosaur. Based on these specimens, Cope named two new species, Ornithochirus umbrosus and Ornithochirus harpyia, in an attempt to assign them to the large European genus Ornithocheirus. As he misspelled the name (forgetting the 'e'), however, he accidentally created an entirely new genus.[5] Cope's paper naming his ''Ornithochirus species was published in 1872, just five days after Marsh's paper. This resulted in a dispute, fought in the published literature, over whose names had priority in what obviously were the same species.[5] Cope conceded in 1875 that Marsh's names did have priority over his, but maintained that Pterodactylus umbrosus was a distinct species (but not genus) from any that Marsh had named previously.[19] Re-evaluation by later scientists has supported Marsh's case, and found that Cope's assertion that P. umbrosus was a larger, distinct species were incorrect.[5]

While the first Pteranodon wing bones were collected by Marsh and Cope in the early 1870s, the first Pteranodon skull was found on May 2, 1876, along the Smoky Hill River in Wallace County (now Logan County), Kansas, USA, by Samuel Wendell Williston, a fossil collector working for Marsh.[1] A second, smaller skull soon was discovered as well. These skulls showed that the North American pterosaurs were different from any European species, in that they lacked teeth. Marsh recognized that this major difference, describing the specimens as "distinguished from all previously known genera of the order Pterosauria by the entire absence of teeth." Marsh recognized that this characteristic warranted a new genus, and he coined the name Pteranodon ("wing without tooth") in 1876. Marsh also reclassified all the previously named North American species from Pterodactylus to Pteranodon, with the larger skull, YPM 1117, referred to the new species Pteranodon longiceps.[20] He also named an additional species, Pteranodon gracilis, based on a wing bone that he mistook for a pelvic bone. He soon realized his mistake, and re-classified that specimen into a separate genus, which he named Nyctosaurus.[21]

Some of the most influential studies of Pteranodon during the 20th Century were published by George Francis Eaton, who conducted thorough re-analysis of the known specimens and published some of the first good photographs and illustrations of the best specimens.[2] During the early 1990s, S. Christopher Bennett also published several major papers reviewing the anatomy, taxonomy and life history of Pteranodon.[4]

Fragmentary fossils assigned to Pteranodon have also been discovered in Skåne, Sweden.[22]

Variation in cranial anatomy and classification of specimens assigned to Pteranodon (drawn to scale, unpreserved portions shown in gray)

Many researchers consider there to be at least two species of Pteranodon. However, aside from the differences between males and females described above, the post-cranial skeletons of Pteranodon show little to no variation between species or specimens, and the bodies and wings of all pteranodonts were essentially identical.[5]

Two species of Pteranodon are traditionally recognized as valid: Pteranodon longiceps, the type species, and Pteranodon sternbergi. The species differ only in the shape of the crest in adult males (described above), and possibly in the angle of certain skull bones.[5] Because well-preserved Pteranodon skull fossils are extremely rare, researchers use stratigraphy (i.e. which rock layer of the geologic formation a fossil is found in) to determine species identity in most cases.

Pteranodon sternbergi is the only known species of Pteranodon with an upright-crest. The lower jaw of P. sternbergi was 1.25 meters (4.1 feet) long.[23] It was collected by George F. Sternberg in 1952 and described by John Christian Harksen in 1966, from the lower portion of the Niobrara Formation. It was older than P. longiceps and is considered by Bennett to be the direct ancestor of the later species.[5]

Because fossils identifiable as P. sternbergi are found exclusively in the lower layers of the Niobrara Formation, and P. longiceps fossils exclusively in the upper layers, a fossil lacking the skull can be identified based on its position in the geologic column (though for many early fossil finds, precise data about its location was not recorded, rendering many fossils unidentifiable).[4]

Below is a cladogram showing the phylogenetic placement of this genus within Pteranodontia from Andres and Myers (2013).[24]

Due to the subtle variations between specimens of pteranodontid from the Niobrara Formation, most researchers have assigned all of them to the single genus Pteranodon, in at least two species (P. longiceps and P. sternbergi) distinguished mainly by the shape of the crest. However, the classification of these two forms has varied from researcher to researcher. In 1972, Halsey Wilkinson Miller published a paper arguing that the various forms of Pteranodon were different enough to be placed in distinct sub-genera. He named these Pteranodon (Occidentalia) occidentalis (for the now-disused species P. occidentalis) and Pteranodon (Sternbergia) sternbergi. However, the name Sternbergia was preoccupied, and in 1978 Miller re-named the species Pteranodon (Geosternbergia) sternbergi, and named a third subgenus/species combination for P. longiceps, as Pteranodon (Longicepia) longiceps. Most prominent pterosaur researchers of the late 20th Century however, including S. Christopher Bennett and Peter Wellnhofer, did not adopt these subgeneric names, and continued to place all pteranodont species into the single genus Pteranodon.

In 2010, pterosaur researcher Alexander Kellner revisited H.W. Miller's classification. Kellner followed Miller's opinion that the differences between the Pteranodon species were great enough to place them into different genera. He placed P. sternbergi into the genus named by Miller, Geosternbergia, along with the Pierre Shale skull specimen which Bennett had previously considered to be a large male P. longiceps. Kellner argued that this specimen's crest, though incompletely preserved, was most similar to Geosternbergia. Because the specimen was millions of years younger than any known Geosternbergia, he assigned it to the new species Geosternbergia maysei. Numerous other pteranodont specimens are known from the same formation and time period, and Kellner suggested they may belong to the same species as G. maysei, but because they lack skulls, he could not confidently identify them.[4]

A number of additional species of Pteranodon have been named since the 1870s, although most now are considered to be junior synonyms of two or three valid species. The best-supported is the type species, P. longiceps, based on the well-preserved specimen including the first-known skull found by S. W. Williston. This individual had a wingspan of 7 m(23 ft).[25] Other valid species include the possibly larger, P. sternbergi, with a wingspan originally estimated at 9 m(30 ft).[25]P. occidentalis, P. velox, P. umbrosus, P. harpyia, and P. comptus are considered to be nomina dubia by Bennett (1994) and others who question their validity. All probably are synonymous with the more well-known species.

P. occidentalis mount wherein arms, shoulder girdle, and fingers are actual bones, and the rest has been drawn from other specimens.

Because the key distinguishing characteristic Marsh noted for Pteranodon was its lack of teeth, any toothless pterosaur jaw fragment, wherever it was found in the world, tended to be attributed to Pteranodon during the late nineteenth and early twentieth centuries. This resulted in a plethora of species and a great deal of confusion. The name became a wastebasket taxon, rather like the dinosaur Megalosaurus, to label any pterosaur remains that could not be distinguished other than by the absence of teeth. Species (often dubious ones now known to be based on sexual variation or juvenile characters) have been reclassified a number of times, and several sub-genera have in the 1970s been erected by Halsey Wilkinson Miller to hold them in various combinations, further confusing the taxonomy (subgenera include Longicepia, Occidentalia, and Geosternbergia). Notable authors who have discussed the various aspects of Pteranodon include Bennett, Padian, Unwin, Kellner, and Wellnhofer. Two species, P. orogensis and P. orientalis, are not pteranodontids and have been renamed Bennettazhia oregonensis and Bogolubovia orientalis respectively.

^ abcdefghijklmBennett, S.C. (1994). "Taxonomy and systematics of the Late Cretaceous pterosaur Pteranodon (Pterosauria, Pterodactyloida)". Occasional Papers of the Natural History Museum, University of Kansas169: 1–70.

Everhart, M. J. 1999. An early occurrence of Pteranodon sternbergi from the Smoky Hill Member (Late Cretaceous) of the Niobrara Chalk in western Kansas. Kansas Academy of Science, Transactions 18(Abstracts):27.

Everhart, M. J. 2005. Oceans of Kansas - A Natural History of the Western Interior Sea. Indiana University Press, 320 pp.

Harksen, J. C. (1966). "Pteranodon sternbergi, a new fossil pterodactyl from the Niobrara Cretaceous of Kansas". Proceedings South Dakota Academy of Science45: 74–77.

Russell, D. A. 1988. A check list of North American marine cretaceous vertebrates Including fresh water fishes, Occasional Paper of the Tyrrell Museum of Palaeontology, (4):57.

Schultze, H.-P., L. Hunt, J. Chorn and A. M. Neuner, 1985. Type and figured specimens of fossil vertebrates in the collection of the University of Kansas Museum of Natural History, Part II. Fossil Amphibians and Reptiles. Miscellaneous Publications of the University of Kansas Museum of Natural History 77:66 pp.

Seeley, Harry G. 1871. Additional evidence of the structure of the head in ornithosaurs from the Cambridge Upper Greensand; being a supplement to "The Ornithosauria." The Annals and Magazine of Natural History, Series 4, 7:20-36, pls. 2-3. (Discovery of toothless pterosaurs in England)